全 文 :书!"#:10004025(2016)08154110 犱狅犻:10.7606/j.issn.10004025.2016.08.1541
%&(
:20160425;)*&%+(:20160612
,-./
:
()*+%,-
(31270241,31470009,31570198,31110103911);(,./012;(34125%34612(ht
tp://mnh.scu.edu.cn);+78,.9:;<=(2013FY112100)
0123
:
>?@
(1990-),A,BCDE:;F,GHIJ#KLMN:;。Email:laistephen@126.com OPQR:ST-,5U,VEFWX,GHIJ#YZ[KLMN\]^_`:;。Email:xingjinhe@aliyun.com
,435,6789:;<犔犲狌犮狅犾犻狉犻狅狀=
>?@ABC5DE?FG
!"#
,
fg[Fhij5k8lmnop
,
qr610064)
H I:stuvw`xFyz{|fg,}u]yz~`KLk[MN]K:;\。:;,
犐犜犛、cpDNA(狋狉狀犞狀犱犺犆、狆犲狋犔狆狊犫犈)
,犃犓犜3,,uY!`yz~6
Z484
GenBank`,yz~犔犲狌犮狅犾犻狉犻狅狀 M¡¢KLk[{£]K:;。¤¥
¦§
:(1)犐犜犛¤¥¨©ª«`¬,®¯犔犲狌犮狅犾犻狉犻狅狀 °Y±2² ,subsect.犔犲狌犮狅犾犻狉犻狅狀6a³subsect.犔犲狌
犮狅犾犻狉犻狅狀6b。´µ¯¶"yz(犔.犺犲狀狉狔犻)³byz(犔.狉狅狊狋犺狅狉狀犻犻)·¸¹subsect.犔犲狌犮狅犾犻狉犻狅狀6a,[º»yz
(犔.狊犪狉犵犲狀狋犻犪犲)、¼½¾yz(犔.狊狌犾狆犺狌狉犲狌犿)、¿Àyz(犔.犾犲狌犮犪狀狋犺狌犿)³ÁÂyz(犔.狉犲犵犪犾犲)4{ÃÄ
q犔犲狌犮狅犾犻狉犻狅狀 ;Å4~犃狉犮犺犲犾犻狉犻狅狀 `xyz(犔.犫狉狅狑狀犻犻)[subsect.犔犲狌犮狅犾犻狉犻狅狀6bƱè,ǯÈ[2
Éyz
(犔.犾狅狀犵犻犳犾狅狉犿)、ÊËyz(犔.犳狅狉犿狅狊犪狀狌犿) qÃÌ ,dͱyz (sect.犔犻犾犻狌犿)。(2) £]K
Î
,犔犲狌犮狅犾犻狉犻狅狀 [yz `KLk]KÏ,ªR[YÐ4`犃狉犮犺犲犾犻狉犻狅狀 ÑqÒÓZ,ÔÕR[犛犻
狀狅犿犪狉狋犪犵狅狀 `Ö]K×。(3),ØÙÚÛz,³,犃犓犜`6{`ÜÝÞßàá,¶
"yz[byz§àâ犔犲狌犮狅犾犻狉犻狅狀 ãtuäåæç¾`4Z。ØÙÚÛz,è³,
褥¦§
,
º»yz
、
¼½¾yz
、
ÁÂyz³¿ÀyzBÜÝé1Φêë¢ìí`\î9
,
ÜÝ]KïY
,
ðñò[yz~#ó‘ôkKLu],õö¢{£ÜÝ÷ø[YN
。
EJK
:犔犲狌犮狅犾犻狉犻狅狀;KLk;犐犜犛;ØÙÚ,;,;{£]K
LMNO :Q789 !PQRS:A 犘犺狔犾狅犵犲狀狔犪狀犱犐狀狋犲狉狊狆犲犮犻犳犻犮犚犲犾犪狋犻狅狀狊犺犻狆狅犳犔犻犾犻狌犿犛犲犮狋犻狅狀 犔犲狌犮狅犾犻狉狅狀犅犪狊犲犱狅狀犜犺狉犲犲犌犲狀犲犛犲狇狌犲狀犮犲狊 LAIShanpan,SHENChengjuan,ZHOUSongdong,HEXingjin (KeyLaboratoryofBioResourcesandEcoEnvironmentofMinistryofEducation,ColegeofLifeScience,SichuanUniversity, Chengdu610064,China) 犃犫狊狋狉犪犮狋:DespiteabundantgermplasmresourcesofwildliliesinChina,thestudyofphylogeneticandinter specificrelationshipin犔犻犾犻狌犿 wasscarce.Inthisstudy,phylogenyofthesection犔犲狌犮狅犾犻狉犻狅狀Comber whichincludesGenBankdataand48specimenscovering6closedrelatedliliesendemictoSouthwestChina wasexploredbasedon犐犜犛,cpDNAandasinglecopynucleargene,犃犓犜.Theresultsshowedthat:(1) inaccordancewithpreviousstudiesthatthesection犔犲狌犮狅犾犻狉犻狅狀Combershouldbedividedintotwosubsec tion,subsect.犔犲狌犮狅犾犻狉犻狅狀6aandsubsect.犔犲狌犮狅犾犻狉犻狅狀6b,and犔.犺犲狀狉狔犻and犔.狉狅狊狋犺狅狉狀犻犻shouldbeclas sifiedintosubsect.犔犲狌犮狅犾犻狉犻狅狀6a.Werevisesection犔犲狌犮狅犾犻狉犻狅狀including犔.狊犪狉犵犲狀狋犻犪犲,犔.狊狌犾狆犺狌狉犲 狌犿,犔.犾犲狌犮犪狀狋犺狌犿,犔.狉犲犵犪犾犲,犔.犺犲狀狉狔犻and犔.狉狅狊狋犺狅狉狀犻犻.犔.犫狅狑狀犻犻,whichoriginalybelongedtosec tion犃狉犮犺犲犾犻狉犻狅狀wasplacedintosubsect.犔犲狌犮狅犾犻狉犻狅狀6b,alongwith犔.犳狅狉犿狅狊犪狀狌犿and犔.犾狅狀犵犻犳犾狅狉狌犿, weestablishanewgroup,namedsection犔犻犾犻狌犿.(2)Asforinfragenericrelationships,section犃狉犮犺犲犾犻狉犻狅狀 whichmainlydistributesinJapanshowedasisterrelationshipwithsection犔犲狌犮狅犾犻狉犻狅狀.However,section 犔犻犾犻狌犿and犛犻狀狅犿犪狉狋犪犵狅狀showedacloserelationship.(3)Thegeneticdistancesamongaltaxaofchloro plastcombinationand犃犓犜showedthat犔.犺犲狀狉狔犻and犔.狉狅狊狋犺狅狉狀犻犻distinctlydifferfromothermembers ofsection犔犲狌犮狅犾犻狉犻狅狀.Phylogenetictreebasedonchloroplastcombinationdata,犃犓犜fragmentindicated that犔.狊犪狉犵犲狀狋犻犪犲,犔.狊狌犾狆犺狌狉犲狌犿,犔.狉犲犵犪犾犲and犔.犾犲狌犮犪狀狋犺狌犿haveaverycloserelationship.Highge neticsimilaritybetweenthesespeciesmaybeattributedtotheparticularmatingsystemof犔犻犾犻狌犿,leading tolimitedgeneticvariationanddifferentiationamongspecies. 犓犲狔狑狅狉犱狊:犔犲狌犮狅犾犻狉犻狅狀;phylogeny;犐犜犛;chloroplastgene;singlecopynucleargene;interspecificrela tionship yz~(犔犻犾犻狌犿 L.)ù~yz+(Liliaceae Juss.),úûüý110~115þ{,ÿ!Y"" #%
,
su55{[1],&yz~#`Äg ³Ys [2], (s)!Y*±vw 。 Comber[3]+13Ñh9ç³2F,9¯y z~-Y±7 9² 。(s,犔犲狌犮狅犾犻狉犻狅狀 `GH.±Ø/F , 01 , ¾±23ç , 6Øë4 56 。 7+89:å;Yq¢2² , Y â ± subsect.犔犲狌犮狅犾犻狉犻狅狀6a ³ subsect. 犔犲狌犮狅犾犻狉犻狅狀6b。ªR<=¢ÁÂyz(犔.狉犲犵犪犾犲)、 ¿Àyz (犔.犾犲狌犮犪狀狋犺狌犿)、º»yz(犔.狊犪狉犵犲狀狋犻 犪犲)³¼½¾yz(犔.狊狌犾狆犺狌狉犲狌犿),89±>å, uYs! ; ÕR?ÊËyz (犔.犾狅狀 犵犻犳犾狅狉狌犿)、2Éyz(犔.犳狅狉犿狅狊犪狀狌犿)³@ABy z (犔.狆犺犻犾犻狆狆犻狀犲狀狊犲) q,89±äå,GHY sC8DE
2É 。 }FG°H%[Y 6KL%:;`I¸ , `-Y
7Z¹ ¢MÃJ`KL[M 。 *N ,Nishikawa[45]
Hayashi[6]yz ~`Y6KL%:;¦§ ,犔犲狌犮狅犾犻狉犻狅狀 O&à K , 7PB2Ö]KÏ`Y¨。Gao [7] yz~`5:;àá , ÑhÎQ±äåæ ç¾`Z , +R£FÚ`u0¯犔犲狌犮狅犾犻狉犻狅狀 Yq2² ,Ss!`ÁÂyz、¿Ày z 、 º»yz³¼½¾yzTuR£FÚUY±¢ ò ; ÔV?yz (犔.狑犲狀狊犺犪狀犲狀狊犲)、WXy z 、 ÊËyz³2ÉyzYZuðä[U·Ã² , ðì]¨©¢Comber[3],89:å `-Y , }\ComberB1949^犔犲狌 犮狅犾犻狉犻狅狀 `-Y,_¸¢`aÕb¦`ÌZ。 ^b , ×ccd7`KLkl´eyz~` YZKLfgh 。Lee[8]yz~83{14÷ {M¡犐犜犛´èYi,犔犲狌犮狅犾犻狉犻狅狀 UYq2 j`Y¨ ,GroupI³ GroupVI。Gao[2,9], 犐犜犛³犿犪狋犓 yz~`KLk:;¤¥àá, ¶"yz (犔.犺犲狀狉狔犻)[subsect.犔犲狌犮狅犾犻狉犻狅狀6aÆ ±Ã¨ , vw¢ðò `{ q
,
k² [Y
Ð4`犃狉犮犺犲犾犻狉犻狅狀 Ñq¢ÒÓZ]K;
subsect.犔犲狌犮狅犾犻狉犻狅狀6bY<=¢xyz(犔.犫狉狅狑
狀犻犻)、WXyz(犔.犪狀犺狌犻犲狀狊犲)、V?yz、ÊËy
z
、
2Éyz³@AByz6{,jƱÃ
¨
,
[subsect.犔犲狌犮狅犾犻狉犻狅狀6a`Ö]KÏ。
Du[10][´¢lª*c`ITSMNè,¤¥¨©
m¶"yz³byz
(犔.狉狅狊狋犺狅狉狀犻犻)·¸¹sub
sect.犔犲狌犮狅犾犻狉犻狅狀6as,犔犲狌犮狅犾犻狉犻狅狀 UKn±2
Y¨
。
)Î`:;¤¥¦§
,犔犲狌犮狅犾犻狉犻狅狀 &O
)*`
,
o(Yp)
°H%
、
Y6KL
%q
,
rHU°Y³s
。
lª¶"yz³byz`KLtuvZu
ûw
,
*NComber[3]+Ñh.¯xR·¸
犛犻狀狅犿犪狉狋犪犵狅狀 ,}Y6KL%`:;[ÑhYZ
¤¥ëê¢yc`zø
,
È{`KLturHlÌ
|}
。
ÔÁÂyz
、
¿Àyz
、
º»yz³¼½¾y
z`ÑhïY\
,
r±äåæç¾
,
}{£ü~
O§à
,
Y6KL%`¤¥àáKLkè`m
¨©Oí
,
Y¨ãPBc`1¡¨
[2,10],
{£]KrHMÃJ:;³
。
4noBª«:
;`,.Î
,
Oc`x³
,
Î
Ss!`Ñh\î
、
Y6KLÎ
`6Z(1),,ØÙÚ,狀犱犺犆狋狉狀犞
2451 ! " # % & 36 A、B.¶"yz³byz(á½åç¾U);C~G.ÁÂyz、¿Àyz、¼½¾yz、º»yz³>yz(áäå 23ç¾U );H、I.¶"yz、byz>å89 1 4:;`yz#Ñh A,B.犔.犺犲狀狉狔犻and犔.狉狅狊狋犺狅狉狀犻犻withorangeTurksCapshapedflowers;C-G.犔.狉犲犵犪犾犲,犔.犾犲狌犮犪狀狋犺狌犿,犔.狊狌犾狆犺狌狉犲狌犿, 犔.狊犪狉犵犲狀狋犻犪犲and犔.犾犲狌犮犪狀狋犺狌犿var.犮犲狀狋犻犳狅犾犻狌犿withwhitetrumpetshapedflowers;H,I.Purplebulbsof犔.犺犲狀狉狔犻and犔.狉狅狊狋犺狅狉狀犻 Fig.1 Imagesof犔犻犾犻狌犿usedinthisstudy ³狆犲狋犔狆狊犫E、Ú,犐犜犛
, 犃犓犜 ,(KLk[{£]KM¡¢Y i:; 。 4:;Z``aq¾c , k7 89ñQ , ÿ!ÜÝk{ , ¾k , Mð{`KLk]K`:;u ±(z`[ ¡w,. 。 1 ¢£³^¤ 1.1 TUVW 4:;I2013~2014^¥Mx)
¦§¨©Q 。 ¶"yz 、 byzGHY lª 、 «¬³¶"Ã% , §¨¹10R。Á yzYÁ® , B¯b 、 °±Ã%§¨¹ 4R。¿Àyz
(÷{>yz犔.犾犲狌犮犪狀 狋犺狌犿var.犮犲狀狋犻犳狅犾犻狌犿 5R,º»yzGH Yab 、 lª , ²11R;Ô¼½¾yzY *±ÿ! , ³ 、 «¬ 、 ÿ!´S , §¨¹ 18R。4:;µ`,¶·)
I GenBank`(¸yz~#`,¶·¹
º
,
犔犲狌犮狅犾犻狉犻狅狀 `KLk[{£]KM¡¢
:;
。
O»¼½7
,
¾R1~2ÚQ±¿
¦
。
¢£À¨PÁ
、
Âq34æ1。ÌÄØ´
ÅÆÇÈÉÊËP
,
Âq34PÌBabc%#
34Í (SZ),¾34`À¨PÁrΧ GPS(GPSMAP62sc,Shanghai)ÏÐÑÒmt。 34518Ó >?@,:,3{,¶·yz~犔犲狌犮狅犾犻狉犻狅狀 `KLk[{£]K:; X1 TUYZ Table1Sourceofmaterials Z Taxon RÔÕ Population code Y Location Öd / ×d Longitude(E)/Latitude(N) Ø5 Haplotype Âq34 Voucherspecimen ¼½¾yz 犔.狊狌犾狆犺狌狉犲狌犿 ZY «¬ÙÚZunyi,Guizhou 106.706006/27.59674 H1 L2014071001 BJ «¬ÛBijie,Guizhou 105.40495/27.59413 H1 L2014070901 GY «¬«ÜGuiyang,Guizhou 106.688921/26.438402 H1 L2014071101 XY «¬TÚXingyi,Guizhou 107.99379/26.57925 H1 L2014071301 XR «¬TÝXingren,Guizhou 105.1571/25.42444 H1 L201407140 AS «¬WÞAnshun,Guizhou 105.9923/26.27725 - L2014071501 DY ³cßDayao,Yunnan 101.09337/25.85903 H19 DY20130630 WSCC ³V? Wenshan,Yunnan 104.111893/23.402378 H19 WS2013070201 WSXC ³V? Wenshan,Yunnan 104.4177778/23.39833333 H4 WS2013070204 WSDT ³V? Wenshan,Yunnan 104.3344444/23.49888889 - WS2013070101 XC ³!àXichou,Yunnan 104.492356/23.260358 - XC20130702 CJ ³áÂChengjiang,Yunnan 102.966077/24.571428 - CJ20130704 JC ³ÂbJiangchuan,Yunnan 102.866741/24.495123 - JC20130704 YM ³âãYimen,Yunnan 102.181305/24.610336 H1(2) YM20130602 ES ³ä?Eshan,Yunnan 102.4516667/24.16777778 H10,H11 ES20130707 JY ³-ÜJinyang,Yunnan 103.255798/27.656657 H1 JY20130731 NJ ³åNanjian,Yunnan 100.316718/24.838849 H1 NJ20130713 HD abæC Huidong,Sichuan 102.631764/26.514499 H22 HD20130717 º»yz 犔.狊犪狉犵犲狀狋犻犪犲 YB ab¿BYibin,Sichuan 102.31653/31.65945 H3,H6 L2014070701 YL ³çèYiliang,Yunnan 104.47764/27.86067 H1 L2014070901 BX abéTBaoxing,Sichuan 102.8175556/30.36263889 H1,H5 L2014080201 EB abäêEbian,Sichuan 103.3372222/29.35583333 H13 EB20130805 MB abëê Mabian,Sichuan 104.0022222/29.35388889 - MB20130804 DJY abrÂìDujiangyan,Sichuan 103.9916667/31.04444444 H12(2) DJY20130626 LD abº»Luding,Sichuan 102.4055556/30.56972222 H1,H17 LD20130728 TQ abíúTianquan,Sichuan 103.2172222/30.11861111 H1,H7 TQ20130729 JFS lª-î?Jinfoshan,Chongqing 107.2197222/29.09111111 H6,H13 JFS20130724 TQLD abíúﺻTianquantoLuding,Sichuan 102.7675/30.02194444 H6(2) 20130826 EMS abäð?Emeishan,Sichuan 103.4211111/29.57472222 H18 EM20130615 ÁÂyz 犔.狉犲犵犪犾犲 WC ab¯b Wenchuan,Sichuan 103.8498/31.70248 H8 L2014062501 LX ab±Lixian,Sichuan 103.84472/31.69307 H8 L2014062502 MX ab°± Maoxian,Sichuan 103.57199/31.46643 H8 L2014062404 HS abñé Heishui,Sichuan 103.3327/31.9828 H2 L2014062801 ¿Àyz 犔.犾犲狌犮犪狀狋犺狌犿 BC ab"bBeichuan,Sichuan 104.16775/31.79763889 H21 L2014080601 QC abòbQingchuan,Sichuan 104.1696389/31.80230556 H20 L2014080701 WX óôV± Wenxian,Gansu 104.7469722/33.24283333 H15 L2014080901 PW ab1õPingwu,Sichuan 104.350575/32.74389 H23 Y2013080501 ZQ óôö÷Zhouqu,Gansu 104.47606/33.798314 - Y2013071502 ¶"yz 犔.犺犲狀狉狔犻 KL «¬øùKaili,Guizhou 107.99389/26.57912 - L2014071201 FL lªúûFuling,Chongqing 107.5684167/29.49375 - L2014081801 YC ¶"¿ÀYichang,Hubei 111.2213611/30.59722222 H9 L2014083101 byz 犔.狉狅狊狋犺狅狉狀犻犻 XE ¶"üýXuanen,Hubei 109.505/29.70722222 - L2014082501 HF ¶"þÿ Hefeng,Hubei 110.0538889/29.98444444 - L20140824 WL lªõ! Wulong,Chongqing 107.9087778/29.54127778 H16 L2014081603 YY lª"ÜYouyang,Chongqing 108.9958333/29.25111111 - L2014082101 ?# 犔.狆狌犿犻犾狌犿 - - H14 Y2012070501
:ZQR±¿Àyz`÷{,>yz
Note:ZQpopulationindicateavarietyof犔.犾犲狌犮犪狀狋犺狌犿,犔.犾犲狌犮犪狀狋犺狌犿var.犮犲狀狋犻犳狅犾犻狌犿
4451 ! " # % & 36 1.2 [ 1.2.1 犇犖犃]V^犘犆犚_` 4:;eío FN+7u~%& (TiangenBiotechCO.LTD., " ) FS`#, DNA ()* 4
:;`µu¦
,
()*¤4|Î&, CTAB
¤
[11],
no®+,()*-§.M¡
。
¹`
DNAS/TE012,ËPB-20℃34s, Ð 。 ÚDNA(nrDNA)` ã5Ò£6(Inter naltranscribedspacers,ITS)&Y6KL%:; *±ÿ!`, 。 4noY6¶·ITSM¡ s78 ( <=ITS1、ITS2)
5.8S),+ White [12] `O9
,
TÚ`78+¶:7
:94℃
;÷92min;s£i±94℃÷945s,45℃<
=45s,72℃>?1min,35@i;*Õ72℃>
?10min。ØÙÚ,狋狉狀犞狀犱犺犆、狆犲狋犔狆狊犫犈
`A°PÁ
78+¶BCShaw[13]。犃犓犜
,9BCDouglas[14],PCR78ÚK[ +¶DQs:7 : <=2.5μL10×PCRbuffer, 200μmol/LdNTPs,2μLDNAEF,2.5mmol/L MgCl2,4μmol/L 9³0.5U犜犪狇DNA G;
95℃;÷94min,s£35@iYâ±95℃÷
930s,58℃>?30s,72℃>?2.5min,*Õ
72℃>?4min。PCR S`HN[I¶©Q? "Jc,+7u~%&Kq , I¶µ9[
PCR9$\Q,nLMNI¶,)ËqI¶¤¥`
OP9
。
1.2.2 abNc I¶¹`MN¶·Laser
geneV0BioEditS0W¸qfasTpV0,B Mega
4[16]U,¶·M¡)Vº³WH`X©s,
sÅYBY Morrison[17]。ZT[¸Q±\]
¬
,
B Mega4sYi¶·`÷øtm(vari
ablesites)、Ë^tm(conservedsites)、_ýPÁt
m
(parsimonyinformationsites)³GC`。À
1000¿a[´bè(neighborjoining,NJ)
:;ZM¡ÆZYi
,kimura2parameter(K2P)
E5²a{£ÜÝÞß
(geneticdistances)。
PAUPver.4.0b10[18]:;Z[´*
c_ýè
(MaximumParsimony,MP),µu`.
Ïu±0¶³O_c
,
Àdpef
(heuristic
search)M¡1000gFhi_(randomaddition)。
)TBR(treebisectionreconnection)M¡jklm
Y¨è
,
²a100n¿,O)M¤(boot
strap,BS)oIpY¨`¨©。Øqè(Bayes
ianinferences,BI)B S 0 MrBayes version
3.1.2[19]Î K q,r s B MrModeltestversion
2.2[20]Îef*zt`E5,P»*tu¿E5±
SYM+G,)Fhè¥,4/ëv+wxyz{
(MarkovChainMonteClarlo,MCMC)Q|¡
10000000,¾1000¿ËP1}è,Ïud± 0.2℃,ª30%`èU~(burnin),7` èzÕÕ¶´èYi 。 2 ¤¥[Yi 2.1 ,678de ,犐犜犛`<`¢98/¶·,< =犔犲狌犮狅犾犻狉犻狅狀 ³犃狉犮犺犲犾犻狉犻狅狀 GHq)
S s`(¸yz~#
,
ºÕ`犐犜犛,¶·
1´kd±621(615~639)bp,<`¢234÷ø
tm
,
PÁtm135,
21.74%,G+C `
62.1%。ØÙÚ,狆犲狋犔狆狊犫犈 ³狀犱犺犆狋狉狀犞
6{`18ÚM¡78I¶,kdYâ
±1192~1196bp³746~753bp,PÁtmYâ
±11³10。?,¶·`PÁtmu~,
狆犲狋犔狆狊犫犈 ³狀犱犺犆狋狉狀犞 M¡¢Ã9kdoo
(ILDoo),¤¥àá/¶·òQRz(犘>
0.5),k±1939~1946bp,` u23÷øt
m
,
PÁtm21(1.1%)。,犃犓犜
:;Z`47ÚM¡78,¶·1´kd±
1459bp,G+C`37.9%,÷øtm100,_
ýPÁtm67(4.6%)。
2.2 fghiNc
)cpDNAÛz,³,犃犓犜 `
¶·±,.
,
MegaS0s`K2PE5
4:;Z`6{3R`18ÚM
¡ÜÝÞßYi
(
¦2)。,cpDNAÛz,`
ÜÝÞߤ¥àá
,6{{£`ÜÝÞ
ß
,
1´ý±0.0044。º»yz、¼½¾yz、
ÁÂyz³¿Àyz`{£ÜÝÞß\î
,
±0.000
~0.002。¶"yz、byz[4äåæç¾Z
`ÜÝÞßB0.006~0.008£。,
犃犓犜B{£`ÜÝÞß1´±0.0199,[ØÙ
Ú,`¤¥Zî
,
ª4{{£ÜÝÞß,c
B0.001~0.007£。¶"yz[byz`
{£ÜÝÞß1´±0.006,[º»yz4{
`ÜÝÞßB0.032~0.036£。
54518Ó >?@,:,3{,¶·yz~犔犲狌犮狅犾犻狉犻狅狀 `KLk[{£]K:;
X2 6jkl5>5Dfghi
Table2 Geneticdistancesamongspeciesin6closerelatedspecies
{Species
º»yz
犔.狊犪狉犵犲狀狋犻犪犲
¼½¾yz
犔.狊狌犾狆犺狌狉犲狌犿
ÁÂyz
犔.狉犲犵犪犾犲
¿Àyz
犔.犾犲狌犮犪狀狋犺狌犿
byz
犔.狉狅狊狋犺狅狉狀犻犻
¶"yz
犔.犺犲狀狉狔犻.
º»yz犔.狊犪狉犵犲狀狋犻犪犲 0.001 0.006 0.001 0.032 0.033
¼½¾yz犔.狊狌犾狆犺狌狉犲狌犿 0.000 0.007 0.001 0.033 0.034
ÁÂyz犔.狉犲犵犪犾犲 0.001 0.001 0.007 0.035 0.036
¿Àyz犔.犾犲狌犮犪狀狋犺狌犿 0.001 0.001 0.001 0.033 0.034
byz犔.狉狅狊狋犺狅狉狀犻犻 0.006 0.006 0.006 0.006 0.006
¶"yz犔.犺犲狀狉狔犻 0.007 0.007 0.007 0.007 0.007
: Î^±,,犃犓犜`ÜÝÞß,7^±cpDNAÛz`ÜÝÞß Note:GeneticdistancesbasedonnuclearsinglecopyAKTareabovediagonal,andgeneticdistancesbasedoncpDNAcombinationarebe lowdiagonal 2.3 ?@ABE?Nc ,犐犜犛¶·[´` MPè³Øqè`¤ ¥àá , {^¤[´`KLkè¤[\î 。 :2µá,犔犲狌犮狅犾犻狉犻狅狀 UY±¢2Y¨,Y â±subsect.犔犲狌犮狅犾犻狉犻狅狀6a³subsect.犔犲狌犮狅犾犻狉 犻狅狀6b,kð2Y¨`KLk]KÏ。sub sect.犔犲狌犮狅犾犻狉犻狅狀6a<`¢º»yz、¼½¾yz、 ÁÂyz³¿Àyz4{,¶"yz³by z[² Ʊè , Y¨[犃狉犮犺犲犾犻狉犻狅狀 Ñq¢ÒÓZ]K (PP/BS,0.85/80)。sub sect.犔犲狌犮狅犾犻狉犻狅狀6b<=¢2Éyz、ÊËyz,x yz[² `ÆZ¤¥¹í`¨© (PP/ BS,0.95/88),sY¨[犛犻狀狅犿犪狉狋犪犵狅狀 `KL k]K×±R 。 B{£]KÎ , 4:;s`6{Ñq¢ Ã`jY¨ ,¨ ©ª«`Y6KL:;¤ ¥ 。 ¶"yz[byzsYNëb , }xR£ `{£]KO§P 。 º»yz 、 ¼½¾yz 、 ÁÂy z³¿Àyz4{`aRZu¹ `ÆZ¤¥ , ã8¦êëî1¡`Y¨ , {£ ü~E , BY6é1Φêë¢ìí`\î9 , } ð4ZBY6KLèλƱÃí¨ ©`KY¨ , ¦§ð4ZtuÃ*` Qs , È{Tu»`Ö]K , ]¨©¢ È{`K9 。 ,ØÙÚ,狀犱犺犆狋狉狀犞 ³狆犲狋犔狆狊犫犈, :;Z`8YÚ ( O<`Z ), ({£ ]KM¡´èYi 。NJèÆZYi¤¥àá( 3),¶"yz³byz`OQRp)Ʊè, tuí`¨© , }xRÑqÃK`KL tu]K 。 Q±äåæç¾`4{Ʊà Bs3¢¨©c50%`¨K;{ÍÕ`
¦áOQRÚÔÕ
3 Ûz2ØÙÚ,(狀犱犺犆狋狉狀犞+狆犲狋犔狆狊犫犈)
6Z[´NJ(O<`)KLkè
Bootstrapvalues>50%aremarked.Theletterplusnumber
behindspecificnamerepresentindividualsfrom
differentpopulations
Fig.3 Phylogeneticrelationshipsofneighborjoining
(nooutgroup)forchloroplastDNAcombination(狀犱犺犆
狋狉狀犞+狆犲狋犔狆狊犫犈)in6closerelatedtaxa
¨
,
¹í`¨©
(99%)。ÁÂyz³¿Ày
z*]p)Ʊè
,
}¨©Oí
,
Bs
Y¨ã8
,
º»yz³¼½¾yz`{£]KOP
»
,
±1¡Y¨lBÃÄ
。
78I¶`ZOÃ
,
kZ`OQ,¶
·] ò
,
µ)ZuOa,Ûz´èbM
ÃJoqyz~ã`KLkYi
。
)?#犔.
狆狌犿犻犾狌犿±Z,,犃犓犜 [´
`KLkè`¤¥àá
(
4),º»yz、¼½¾
yz
、
ÁÂyz³¿Àyz4{Ʊ¢Ã
¨
(PP/BS,0.98/97),}È{`{£]K¡*O§
P
,
PBÃc`1¡Y¨
。
¶"yz
、
byz
[Y¨`KL]K¢
,
}Zu¨©
。
6451 ! " # % & 36 m¬Y⿦ØqèÕo£ (PP)³*c_ýèBootstrap¨©(BS),{ÍÕ¤[Yâ¦áGenBank Õ³RÚÔÕ 。 7Q 2 ,犐犜犛¶·犔犲狌犮狅犾犻狉犻狅狀
\]Z`KLkYi Bayesianposteriorprobabilities(PP)andparsimonybootstrap(BS)aregivenatnodes,respectively Theletterplusnumberbackofspecificnamerepresentpopulationorindividualcodes.Thesameasbelow Fig.2 Phylogenyanalysisofsection犔犲狌犮狅犾犻狉犻狅狀anditsrelatedtaxabasedon犐犜犛sequence 74518Ó >?@,:,3{,¶·yz~犔犲狌犮狅犾犻狉犻狅狀 `KLk[{£]K:; 4 ,,犃犓犜6 Z`KLkYi Fig.4 Phylogenyanalysisof6closerelatedtaxa basedonthenuclearsinglecopysequence犃犓犜 Ø5Yi¤¥àá ,42/,¶·ê¢23{ Ø5 ( ¦1),IØ5¥¦Î§(5),Ø5H1 &Y*ÿ , ëê¨*í`Ø5 , g)º»yz ³¼½¾yz`c8YR , ©Ø5 H6、H13、 H19]r&b)º»yz³¼½¾yzR,ª H8«,úrb)ÁÂyzR。¬uØ5v w ,H2、H3、H4、H5、H7、H15、H17、H18、H21、 H23[Ø5 H1tu,¦êë¢ì±Ö` tu]K 。 3
w 3.1 犔犲狌犮狅犾犻狉犻狅狀=>?@ABNc ª«Bûü®¯ãsyz~`KLk¹ ¢±ú_`:; , lª`:;¦§ , B)Y63Ñ µ[´`yz~KLkès , syz~ (` °6 ¾~ ) [qñ)*`KZ ,Comber[3], Ñh.´7 `yz~~7YZKL¹ ¢ca%R`±ñ 。 *Ô , Y6KLk:;¦ § , yz~`Y6KL[Comber,Ñh.´ `yz~`YZKL£PBc`zø ,Comber ´`yz~~77 ²³[Y6Zu´c`] Û9 。 ª«:;¤¥¦§ , ¶"yz[犔犲狌犮狅犾犻狉犻狅狀 µ¶c·êë¾Ã{Ø5`¨ ; Z`µm¿¦ s£Ø5 , B4:;s oI¹¸¹Öº» 5 ,,犃犓犜6{[´` Ø5¥¦]K Thesizeofthecirclesindicatesthefrequencyofeachhaplotype. Emptydotsindicatethatintermediatehaplotypeswerenot detectedinourresearchorextinction Fig.5 Relationshipswthinthehaplotypenetwork for6closerelatedtaxa ¦êë¢ì±`]K [45], Ôbyz`\] :;¼
。 4:;§¨6{`aR ¦ , Ûz\]Zyz~M¡KLkl´ 。 犐犜犛褥àá, -Yë¢2jY¨,T uäåæç¾`º»yz 、 ¼½¾yz 、 ¿Àyz³ ÁÂyz , [tu½åL¾U`¶"yz 、 byzY6KL]Kì±Ö 。 ðä¥[ª«: ;¤¥Ã , ¶"yz[byz½·¸¹sub sect.犔犲狌犮狅犾犻狉犻狅狀6as,kxRrtu>å89ð Ã. , MÃJ¾q¢Comber[3],89:å 犔犲狌犮狅犾犻狉犻狅狀 ` 7-Y。ÔÅ4~犃狉犮犺犲犾犻狉犻 狅狀 `xyz[subsect.犔犲狌犮狅犾犻狉犻狅狀6bƱà ¨ , BKLtuÎ[犛犻狀狅犿犪狉狋犪犵狅狀 `Ö]K ×±R 。 + 《 s#¿
》[21]
`ÑhYZ¬
,
¤zY6KLk]K
,
犔犲狌犮狅犾犻狉犻狅狀 M¡¢
lÌ-Ys
。
QÀí³C
[22]
` µ
,¯
º»y
z
、
¼½¾yz
、
¿Àyz
、
ÁÂyz)
Ì-¸`
¶"yz³byz6{ q犔犲狌犮狅犾犻狉犻
狅狀 ,È{´±s!u{,ð`¬
]Áz `Ñh.
(
89>å
)
³YE
p
。
ª«`{£Âl]¦§¢¶"yz[º»y
z
、
¼½¾yz
、
¿Àyz
、
ÁÂyztu`Ö
]K
[2325]。
¯xyz[subsect.犔犲狌犮狅犾犻狉犻狅狀6b
s`2Éyz
、
ÊËyz qÃÌ
,
dͱy
z
(sect.犔犻犾犻狌犿),(qcaYBC²DE
8451 ! " # % & 36 ,´ Tu23ç¾Ã , äå89. 。 I £]KΧ ,犔犲狌犮狅犾犻狉犻狅狀 [YBC² DE`犃狉犮犺犲犾犻狉犻狅狀 ÑqÒÓZ,Ôyz BKLtuÎ[犛犻狀狅犿犪狉狋犪犵狅狀 ×±R[2,10],ð à £]KB{£Âl(o]¹¢¨© 。 O» , Ö]K×`{
,
ÂlqÄ×í
。
ÅÆÎ`
C^yzÂlKÇ&I犃狉犮犺犲犾犻狉犻狅狀 ã`犔.
狊狆犲犮犻狅狊狌犿、犔.犼犪狆狅狀犻犮狌犿 [¶"yz`Âl{s
kÔb
。
IÜÝ%dÎ-
,
Y6KLYZ`¤
¥×_OPñ¢
,
×_»
。
Ô,ÑhµQë`
YZ¤¥Y+YZ.ÈlmOQÔ¦êë
c`zø
。
3.2 klOm>5DE?
4:;(µ`6{`{£KL
]KM¡MÃJ`P±
。
IØÙÚÛz,³
,犃犓犜 ²a¹`{£ÜÝÞßb§,¶
"yz³byz*[º»yz
、
¼½¾yz
、
Á
Âyz
、
¿ÀyzƱè
,
}OæBÑhÎv&Ü
ÝÞßÎr[Õ4{zø§à。,¶"yz
³byz`{£ü~E
,
xR`ÑhïY\î
,
rTu½åL¾U
、
>å89³5Ø
.
;
ðxR`{£]Ku&É
,Du[10]
ÊËÅtÂl7ÌBÍåÚé1Î-§¢R`
âBBÍåÚ`u0Î,*ÔBÍåÚðä[
OòQ±OQ{`YZ+
,
±È`PB[Î
×a[ij/0\]
[22,26]。
¶"yzGHY
«¬
、
lª³¶"jã`õû??Ï
Ð??Ï`
ÑÒFjÎ
,
RYTÓ\6ß
,
Ôbyzª
êlª³¶"lü¬`ðԮ`ÕÖ¸
R×Øs
,
DÔ®FkY
,
xRBFjÙÎu
c`zø
。
4:;,犐犜犛[´`KLkè
àá
,
*xRòƱÃj`Y¨
,
}IaR
`¤¥Oò_)Y
。
BØÙÚÛ
z,ès
,
b)OQR`2{`Ú
p)ÆZBÃY¨ã
,
}BKLtu]KÎ&Ã
KY¨
,
-§¢xRBÜÝé1ÎPBGÿ!
`l®
。
ÑhÎ
,
¶"yz[byzGHâB
Ø[¥n`c
;
*Ô
,
Y6é1ÎxR¦ê
ë¢ìí`\î9
,
ÜÝÞß
,
]ð2{`
{£ü~vrHMÃJ
w
。
Q±äåæç¾`º»yz
、
¼½¾yz
、
ÁÂ
yz
、
¿ÀyzBÑhÎïY\
,
}KLtuÃÚ
\»
。
ØÙÚÛz,è òÛÜÈ
{£`]K
,4ZƱÃc`1¡Y¨,Ý
±ÒÓZ]K
,
kTuí`¨©
。
ð-§
¢ð4Z`K9,Ö]KÞ。
,` MPè³Ø5Yi]QOòYð4
Z
。
IYΧ
,
4Z`Y
,
¾ÓßïY\î , ±{£`)*Âl ¡¢ñ ò , àM¢È{`,l® , IÔB,é1¦êë á`\î9 , ðÃmByz+â3¾~Zs ]PB [27]。 âá`&º»yz³¼½¾yz , ðxRIYΧ*±ÿ! , kZu§à ã6 , Ñhzøì , ròSFä6M¡09ô, , ð_È¢xR`7å[Ýæ , Bª«:;&És , Ö »mxRçè , µ)¸{`{£]KvrHMÃJ
。 4:;`6Z`{ãÜÝ÷ø\z , B,é1ÎÜÝzøO§à 。 R£`ÜÝ YN³Üݤ[éé[,® 、 ¾ê{6`Ýæ)
ôkKLë\] [28]。 yz~#tu±
ìÂÔ;íú`ô,hî
,
ò)89M¡09F,
,
ðeu9F,+ï
,
±yz`,ðËP
¡¢Ëñ
,
}Q|]Ì0¢,÷ø[YN`È
,
eyz~#B,é1ηêë¢ìí`\î 9 , {£`ÜÝzøO§à , ò{£]K`ó»Îô q¢ìc`õö 。 ÷ε , 4:;,3{Y6¶·Yi¢y z~7 犔犲狌犮狅犾犻狉犻狅狀 `KLk]K,§P¢ 犔犲狌犮狅犾犻狉犻狅狀 PB2j`Y¨,¯ ¶"yz、 byz 、 º»yz 、 ¼½¾yz 、 ÁÂyz³¿À yz6{·¸¹Qà s,[q犔犲狌 犮狅犾犻狉犻狅狀 。( ã6Z`{£]K¡O §P , rHøù×aY6XM¡×±I¸`:; 。 no!P : [1] LIANGSY,TAMURA M N.FloraofChina[M].Beijing: SciencePress,2000,24:135149. [2] GAOYD,HARRISAJ,ZHOUSD,犲狋犪犾.Evolutionarye ventsin犔犻犾犻狌犿 (including犖狅犿狅犮犺犪狉犻狊,Liliaceae)aretempo ralycorrelatedwithorogeniesoftheQTplateauandthe HengduanMountains[J].犕狅犾犲犮狌犾犪狉犘犺狔犾狅犵犲狀犲狋犻犮狊犪狀犱犈狏狅 犾狌狋犻狅狀,2013,68(3):443460. [3] COMBERHF.Anewclassificationofgenus犔犻犾犻狌犿[J].犔犻犾狔 94518Ó >?@,:,3{,¶·yz~犔犲狌犮狅犾犻狉犻狅狀 `KLk[{£]K:; 犢犲犪狉犫狅狅犽,犚狅狔犪犾犎狅狉狋犻犮狌犾狋狌狉犪犾犛狅犮犻犲狋狔 (犔狅狀犱狅狀),1949,15: 86105. [4] NISHIKAWAT,OKAZAKIK,UCHINOT,犲狋犪犾,NAGA MINET.A molecularphylogenyof犔犻犾犻狌犿intheinternal transcribedspacerregionofnuclearribosomalDNA[J].犑狅狌狉 狀犪犾狅犳犕狅犾犲犮狌犾犪狉犈狏狅犾狌狋犻狅狀,1999,49:238249. [5] NISHIKAWAT,OKAZAKIK,ARAKAWAK,犲狋犪犾.Phy logeneticanalysisofsection犛犻狀狅犿犪狉狋犪犵狅狀ingenus犔犻犾犻狌犿u singsequencesoftheinternaltranscribedspacerregioninnu clearribosomalDNA[J].犅狉犲犲犱犻狀犵犛犮犻犲狀犮犲,2001,51(1):39 46. [6] HAYASHIK,KAWANOS.Molecularsystematicsof犔犻犾犻狌犿 andaliedgenera(Liliaceae):phylogeneticrelationshipsamong 犔犻犾犻狌犿andrelatedgenerabasedonthe狉犫犮犔and犿犪狋犓gene sequencedata[J].犘犾犪狀狋犛狆犲犮犻犲狊犅犻狅犾狅犵狔,2000,15(1): 7393. [7] GAOYD,ZHOUSD,HEXJ.Karyotypestudiesinthirty twospeciesof犔犻犾犻狌犿 (Liliaceae)from China[J].犖狅狉犱犻犮 犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔,2011,29(6):746761. [8] LEECS,KIMSC,YEAUSH,犲狋犪犾.Majorlineagesofthe genus犔犻犾犻狌犿 (Liliaceae)basedonnrDNAsequences,with specialemphasisontheKoreanspecies[J].犑狅狌狉狀犪犾狅犳犘犾犪狀狋 犅犻狅犾狅犵狔,2011,54(3):159171. [9] GAOYD,HOHENEGGER M,HARRISA,犲狋犪犾.Anew speciesinthegenus犖狅犿狅犮犺犪狉犻狊Franchet(Liliaceae):evi dencethatbringsthegenus犖狅犿狅犮犺犪狉犻狊into犔犻犾犻狌犿[J].犘犾犪狀狋 犛狔狊狋犲犿犪狋犻犮狊犪狀犱犈狏狅犾狌狋犻狅狀,2012,298:6985. [10] DUYP,HEHB,WANGZX,犲狋犪犾.Molecularphylogeny andgeneticvariationinthegenus犔犻犾犻狌犿 nativetoChina basedontheinternaltranscribedspacersequencesofnuclear ribosomalDNA[J].犑狅狌狉狀犪犾狅犳犘犾犪狀狋犚犲狊犲犪狉犮犺,2014,127 (2):249263. [11] DOYLEJ.ArapidDNAisolationprocedureforsmalquanti tiesoffreshleaftissue[J].犘犺狔狋狅犮犺犲犿.犅狌犾犾.,1987,19: 1115. [12] WHITETJ,BRUNST,LEES,犲狋犪犾.Amplificationsand directsequencingoffungalribosomalRNAgenesforphyloge netics[M]//INNISMA,GELFANDDH,SNINSKYJJ. PCRProtocols:AGuidetoMethodsandApplications.San Diego:AcademicPress,1990:315322. [13] SHAWJ,LICKEYEB,SCHILLINGEE,犲狋犪犾.Compari sonofwholechloroplastgenomesequencestochoosenoncod ingregionsforphylogeneticstudiesinangiosperms:thetor toiseandthehareIII[J].犃犿犲狉犻犮犪狀犑狅狌狉狀犪犾狅犳犅狅狋犪狀狔, 2007,94(3):275288. [14] DOUGLASNA,WALLWA,XIANGQY,犲狋犪犾.Recent vicarianceandtheoriginoftherare,edaphicalyspecialized Sandhilslily,犔犻犾犻狌犿狆狔狉狅狆犺犻犾狌犿 (Liliaceae):evidencefrom phylogeneticandcoalescentanalyses[J].犅犿犮犆狅犿狆犾犲犿犲狀狋犪 狉狔牔犃犾狋犲狉狀犪狋犻狏犲犕犲犱犻犮犻狀犲,2011,20(14):29012915. [15] BURLANDTG.DNASTARsLasergenesequenceanalysis software[J].犕犲狋犺狅犱狊犻狀犕狅犾犲犮狌犾犪狉犅犻狅犾狅犵狔,2000,132:71 91. [16] TAMURAK,DUDLEYJ,NEIM,犲狋犪犾.MEGA4:Molec ularevolutionarygeneticsanalysis(MEGA)softwareversion 4.0[J].犕狅犾犲犮狌犾犪狉犅犻狅犾狅犵狔 牔 犈狏狅犾狌狋犻狅狀,2007,24(8):1 5961599. [17] MORRISONDA.Aframeworkforphylogeneticsequencea lignment[J].犘犾犪狀狋犛狔狊狋犲犿犪狋犻犮狊牔 犈狏狅犾狌狋犻狅狀,2009,282 (3):127149. [18] SWOFFORDDL.PAUP4.0:PhylogeneticAnalysisU singParsimony(andOtherMethods)[M].Sunderland: SinauerAssociates,2001. [19] FREDRIKR,HUELSENBECKJP.MrBayes3:Bayesian phylogeneticinferenceundermixedmodels[J].犅犻狅犻狀犳狅狉犿犪狋 犻犮狊,2003,19(12):15721574. [20] NYLANDERJ.MrModeltest2.0:Programdistributedby theauthor[M].EvolutionaryBiologyCentre,UppsalaUni versity,2004. [21] úd,` ûð,üýþ.s#¿[M].":+%ëFÿ,
1980:116.
[22] í³C.yz~
(~#`5MN
yz~`Y6 KL% 、 F%:;
[D].qr:abc%,2013.
[23] LIGHT R W.Cytologicalandinterspecifichybridization
studiesin犔犻犾犻狌犿 L.andtheirsignificanceforclassification
[D].Ithaca:CornelUniversity,1960.
[24] SYNCEPM.Lilies[M].London:Batsford,1980.
[25] WOODCOCK H BD,STEARN WT.Liliesoftheworld
[M].London/NewYork:CountryLife/Scribner,1950.
[26] !"#.#°HYZ%[M].":+%ëFÿ,1990. [27] PETERSON A,JOHN H,KOCH E,犲狋犪犾.A molecular phylogenyofthegenus犌犪犵犲犪 (Liliaceae)inGermanyin ferredfrom noncodingchloroplastandnuclearDNA se quences[J].犘犾犪狀狋犛狔狊狋犲犿犪狋犻犮狊牔犈狏狅犾狌狋犻狅狀,2004,245(3): 145162. [28] PETERSON A,BARTISHIV,PETERSONJ.Genetic structuredetectedinasmal populationoftheendangered plant犃狀狋犺犲狉犻犮狌犿犾犻犾犻犪犵狅(Anthericaceae)byRAPDanalysis [J].犈犮狅犵狉犪狆犺狔,2002,25(6):677684. ( !" : #%
)
0551 ! " # $ % & 36